Volume 22, Issue 4, Pages 751-761 (April 1999) Increased Activity in Human Visual Cortex during Directed Attention in the Absence of Visual Stimulation  Sabine Kastner, Mark A. Pinsk, Peter De Weerd, Robert Desimone, Leslie G. Ungerleider  Neuron  Volume 22, Issue 4, Pages 751-761 (April 1999) DOI: 10.1016/S0896-6273(00)80734-5

Figure 1 Experimental Design Four complex images were presented in four nearby locations at 6°–10° eccentricity from a fixation point (FP) either sequentially (A) or simultaneously (B). Presentation time was 250 ms, followed by a blank period of 750 ms, on average, in each location. A stimulation period of 1 s is shown, which was repeated in blocks of 10 s interleaved with blank periods (BL, shown in [C]). Two attentional conditions were tested (C): an unattended (UNATT), during which subjects maintained fixation, and an attended (ATT), during which subjects covertly directed attention to the location closest to fixation and counted the occurrences of one of the images. An attended condition was indicated 10 s before the onset of visual presentations. During this period (EXP), subjects covertly directed attention to the target location expecting the occurrences of the presentations. Blocks with expectation and attended presentations could be either the first and third, as shown in (C), or the second and fourth with the sequence counterbalanced across scans. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)

Figure 2 Activated Areas in Visual Cortex (A) Areas in ventral visual cortex activated by the complex images in the unattended condition compared to the subsequent blank presentations (control) plotted onto coronal slices of a single subject at different distances from the posterior pole. Activated voxels were assigned to areas V1, V2, VP, V4, and TEO as determined by retinotopical mapping. (B) Activated voxels in the same subject and the same slices contrasting the expectation period with the respective blank period preceding an unattended presentation (control). Areas V2, V4, and TEO showed significant clusters of activation. R, right hemisphere. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)

Figure 3 Time Series of fMRI Signals in Visual Cortex Time series of fMRI signals in V1, V2, V4, and TEO averaged over all subjects. Gray shades indicate the expectation period, blue shades the attended presentations. Blocks with unattended presentations are shown without shading. During the unattended presentations, sequentially presented stimuli evoked stronger activity than simultaneously presented stimuli. This sensory suppression effect was strongest in V4 and TEO. In the attended condition, signals to simultaneously presented stimuli increased to a larger extent than to sequentially presented stimuli. When subjects directed attention to the target location during the expectation period, activity increased in the absence of visual presentations and further increased after the onset of visual stimuli. This baseline increase was seen in all visual areas under investigation, with the strongest effect in V4. SEQ, sequentially presented stimuli; SIM, simultaneously presented stimuli. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)

Figure 4 Mean Signal Changes and Indices of Attentional Modulation and Baseline Shift in Visual Cortex (A) Mean signal changes in V1, V2, V4, and TEO, averaged across subjects. For each subject, the five peak intensities of the fMRI signal obtained during attended and unattended presentations and during the expectation period were averaged. Activity during the expectation period (EXP) is shown compared to two different control conditions: in the first, used in the main experimental design, subjects simply maintained fixation (EXPSF; SF, simple fixation); in the second, used in the control experiment, subjects counted Ts or Ls at fixation (EXPT/L). (B) Attentional response modulation on sequentially and simultaneously presented stimuli as quantified with an AMI, which normalizes the attentional effects to the activity evoked in the respective attended condition. Similarly, the BSI indicates the amount of baseline increase relative to the activity evoked by the subsequent attended presentations. BSIs are only shown for data obtained with the main experimental design. In V2, V4, and TEO, the baseline increases were similar to the attentional effects on simultaneous presentations but stronger than the attentional effects on sequential presentations. Vertical bars indicate SEM. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)

Figure 5 Areas in Frontal and Parietal Cortex Activated during Expectation Same subject as in Figure 2. (A) Activated areas in frontal (SEF, FEF) and parietal (SPL, IPL) cortex obtained in session 2 and depicted on axial slices. The approximate Talairach plane is indicated for each slice. (B) Activated areas in parietal cortex (SPL, IPS) obtained in session 1. Same coronal slices as shown in Figure 2. The distances from the posterior pole are indicated. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)

Figure 6 Time Series of fMRI Signals in Frontal and Parietal Cortex Time series of fMRI signals in IPS, SPL, FEF, and SEF averaged across subjects. Conventions as in Figure 3. During the expectation period, signals increased to a greater degree in FEF, SEF, and SPL than in visual cortex; this response increase was not followed by a further increase after the onset of the stimuli resulting in a sustained activity. The pattern of activity in IPS resembled more that in extrastriate areas than in SPL, FEF, and SEF. None of the frontal or parietal areas showed sensory suppressive effects among interacting visual stimuli. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)

Figure 7 Mean Signal Changes and Indices of Attentional Modulation and Baseline Shift in Frontal and Parietal Cortex All conventions as in Figure 4. All areas showed significant attentional response enhancement. Attentional modulation on sequentially and simultaneously presented stimuli was similar in all areas. Baseline increases were stronger than attentional modulation effects in all areas. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)

Figure 8 The Relation of Attentional Modulation and Increases in Baseline Activity in Frontal, Parietal, and Visual Cortex For each area, the BSI is plotted against the mean AMI, averaged across sequential and simultaneous conditions. Attentional modulation and baseline increases were similar in ventral extrastriate areas V2, V4, and TEO and parietal area IPS. In V1, a relatively strong increase of baseline activity was seen, whereas attentional modulation was weak. SPL, FEF, and SEF showed a similar relation of attentional modulation and baseline increases. Both effects were, however, stronger than in extrastriate cortex and IPS. Neuron 1999 22, 751-761DOI: (10.1016/S0896-6273(00)80734-5)